Chapter 1

Vertebrata

1.1 Introduction
The Phylum Chordata comprises three sub-phyla, namely, Urochordata, Cephalo-
chordata and Vertebrata. The notochord, a post-natal tail, a dorsal nerve cord
(tube) and pharyngeal gill slits are the four major characteristics of Chordata, and
are present in the species either all through or partially (some time) in the species
comprising this phylum.

Sub-phylum Vertebrata, has the following characteristics:

• The notochord is embryonic (in most cases) and is replaced by a vertebral

column - except in "Agnathans"

• The anterior part of the nerve cord becomes differentiated into a brain - which
means the organisms have a distinctive head or cephalisation has occurred

• Closely connected to the brain are the - nose, eye and ear - a specialized and
highly advanced sensory system

• Surrounding the brain and the sense organs are the cranium (acranium)

• Gill slits are usually five - exists only in the embryonic stages (reptiles, birds
and mammals, exists throughout life (fishes) or in larval stages (amphibia)

• Growing endoskeleton (bones or cartilage).

• Embedded in the pharynx (between the gill slits) are the visceral skeleton
(to support the viscera)

• Paired appendages are present - always two pairs (or less in number) -
agnathans do not have paired limbs

• Paired mesodermal somites are present in the embryo - which is metameric

segmentation, metameric segmentation is also evident in adults in the ar-
rangement of vertebrae, muscles and nerves

• Chambered ventral heart as the centre of the circulatory system is a key

vertebrate characteristic

1
2 1.2. CLASSIFICATION AND GENERAL CHARACTERISTICS

• A complete digestive tract

• The presence of an endostyle or a thyroid gland is another distinctive char-
acteristic
Looking at the characteristics above we readily acknowledge that the group de-
serves a better name, since the agnathans are also included with this group, some
zoologists prefer the name "Craniata", for this whole group and vertebrata for the
group excluding agnathans.

1.2 Classification and general characteristics

Vertebrates can be broadly classified as follows:

Table 1.1: Classification of Vertebrata (Craniata)

Sub-phylum Craniata (Vertebrata)

Division I: Agnatha
Class I: Myxini (eg: hagfishes)
Class II: Ostracodermi (Extinct)
Class III: Cephalaspidomorphi (eg: lam-
preys)
Division II: Gnathostomata
Super-class 1: Pisces
Class I: Placodermi (Extinct)

Class II:Chondrichthyes
(Sub-class 1: Elasmobranchii eg: sharks,
rays and skates)
(Sub-class 2: Holocephali eg: ratfishes or
chimeras)

Class III: Osteichthyes

(Sub-class 1: Actinopterygii eg: ray-
finned fishes)
(Sub-class 2: Sarcopterygii eg: lobe-
finned fishes)

Super-class 2: Tetrapoda
Class I:Amphibia
Class II:Reptilia
Class III:Aves
Class IV:Mammalia

“Agnathans” - Lack jaws and paired appendages they possess a cartilaginous

skeleton and have a persistent notochord (ie., they do not have vertebra) two semi-
circular canals. (Hagfishes have one semicircular canal that may represent a fusion
of two canals).

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CHAPTER 1. VERTEBRATA 3

Class Myxini comprises the hagfishes- possess a mouth with four pairs of ten-
tacles they have their olfactory sacs opening to their mouth cavity and have 5 to 15
pairs of pharyngeal slits. They live in mud (in the bottom) of marine environments
and feed on invertebrates and scavenge on dead and dying fishes. Most zoologists
consider hagfishes to be the primitive vertebrate.
Class Cephalaspidomorphi comprise the Lampreys - they have a sucking mouth
with teeth and rasping tongue, have seven pairs of pharyngeal slits and blind olfac-
tory sacs. Lampreys are found in marine and freshwater environments of temperate
regions. Their larvae are filter feeders and the adults prey on fishes. Mouth is suck-
erlike and surrounded by lips for attachment and sensory functions. They live in the
oceans and migrate to freshwaters for spawning. They lay eggs, which develop into
ammocete larvae which live in the mud (bottom) filter feeding for almost 3 years.
The larvae mature into adult and migrate back to the sea to lead a predatory adult
life.
“Gnathostomes” (Division Gnathostomata) have hinged jaws and paired ap-
pendages. They are more properly called as jawed vertebrates. They have a vertebral
column and in some cases notochord is persistent in between some vertebrae, gen-
erally the vertebrae (bones) replace the notochord these organisms possess three
semicircular canals. The major innovations during vertebrate evolution was the ap-
pearance of ’jaws and paired appendages’. Gnathostomes possess both these, which
allows them to live a more active and predatory lifestyle.
There are two major classess of organisms belonging to the ’gnathostomata’
which are still alive (extant). The chondrichthyes (cartilaginous fishes) and the
osteichthyes (bony fishes).

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4 1.2. CLASSIFICATION AND GENERAL CHARACTERISTICS

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Chapter 2

Super-class Pisces

2.1 introduction
What is a fish?
Berra (2001) stated that a fish is a -

• a poikilothermic, aquatic chordate with appendages (when present) developed

as fins, whose chief respiratory organs are gills and whose body is usually
covered with scales

Nelson in 2006, provides a simpler definition -

• a fish is an aquatic vertebrate with gills and with limbs in the shape of fins

Normally defining a fish is a difficult process, since exceptions are quite often.
In addition every organism in Agnatha and Gnathostomata are called as fishes.
The hagfishes, lampreys, sharks, rays, skates, ratfishes, ray-finned fishes, lobefinned
fishes (lungfishes), gars, sturgeons, paddlefishes etc. are all called as fishes. How-
ever, as students of zoology we know that Agnathans have to be excluded when we
strictly speak about fishes.
Fishes (super-class: Pisces) are of two classes (or groups in lay terms), the
cartilagenous fishes and the bony fishes.
Chondrichthyes, (In Greek: chondros - cartilage, ichthyos - fish), include sharks,
rays, skates and ratfishes. They have a tough skin, with placoid scales. Most of
the species have a caudal fin with the upper lobe being large (heterocercal caudal
fin). They possess biting mouth parts and paired appendages (fins), and possess a
cartilaginous endoskeleton.
The sub-class elasmobranchi (sharks, rays and skates) have around 700 species.
The sharks are though to have arose (during evolution) in the Devonian period
around 375 million years ago.
The placoid scales, are pointed backwards, on the skin gives it a sandpaper
texture. The fact that the scales are pointed backwards, nullifies any resistance
(reduce friction) while swimming. The shark teeth, are modified placoid scales and
are present as a row on the outer edge of the jaw, and inner row of teeth are present
on a ligamentous band covering the jaw. The teeth on the outer edge are replaced
by the inner (younger) ones periodically (some 7-8 days in young sharks. Sharks
range in size from less than 1 metre to 10 meters. The largest sharks (whale sharks

5
6 2.2. SCOLIODON SORRAKOWAH

and basking sharks) are not predatory but are filter feeders, with their pharyngeal
arches modified for filter feeding.
Skates and rays, are specialized to live in the ocean floor in shallow waters.
They use their blunt teeth to feed on marine invertebrates. They have wing like
extensions on the lateral side of their body, which are actually modifications of their
pectoral fins. The sting ray (Dasyatis) has a tail modified into a defensive lash; the
dorsal fin persists as a venomous spine. Also included in this group are the electric
rays (Narcine and Torpedo) and manta rays (Manta).
The second sub-class of cartilaginous fishes are the Holocephali (in Greek: Holos
- whole, kephalidos - head), which are normally known as chimeras. The chimeras
(or holocephalans) lack scales. They have large cutting plate like teeth to crush
marine invertebrates. They have evolved to have a covering over their gill slits -
operculum - which is a norm in bony fishes, but not seen in other cartilaginous
fishes. They have a large head, small mouth with large lips, and a narrow tapering
tail (which is why they are called as rat fishes).

2.2 Scoliodon sorrakowah

Scoliodon is a common requiem shark in the indo-pacific. [Requiem sharks are
a family, Carcharhinidae, of sharks in the order Carcharhiniformes, containing
migratory, live-bearing sharks of warm seas (sometimes of brackish or fresh water)
such as the tiger shark, the blue shark, the bull shark, and the milk shark].

2.3 General morphology

This shark grows upto 2 feet, has a long spindle-shaped body (the body shape offers
no resistance for swimming since the body tapers at both ends). The body can be
divisible into a head, trunk and a tail even though there is no clear demarcation of
these three parts. Head is depressed and produced into a snout, trunk is elliptical
and slightly flattened from side to side, the tail is narrow and compressed laterally.
Body is dark-grey in colour above (dorsal side) and slightly paler on the ventral side.

2.3.1 Placoid scales

The scales on the body of these sharks are placoid (in Greek: plax - plate, eidos -
form), formed on a basal plate. They have tri-radiate spines, with their (the spines’)
ends pointing backwards (giving it a sandpaper texture). The outer covering is of
enamel and internal to it is dentine, with a pulp cavity in the interior (the pulp
cavity contains pulp which encloses the dentine-forming cells or odontoblasts).
The basal plate is made up of calcified tissue, it also has an opening through which
blood vessels and nerves enter the pulp cavity.
Placoid scales are formed when a mass of cells aggregate in the dermis, im-
mediately below the epidermis. These cells bulge to form a dermal papilla. The
epidermal cells lining the papilla above function as the enamel organ, the outermost
layer of dermal cells in the papilla function as odontoblasts and the bulge grows,
finally erupting out through the epidermis as spines pointed backwards. The enamel
at the base of the bulge in the meantime forms the basal plate and the whole scale

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CHAPTER 2. SUPER-CLASS PISCES 7

is innervated and supplied with blood vessels through a small opening in the basal
plate.

2.3.2 Fins
Fins are characteristic of fishes. They are flattened expansions of the skin supported
by skeletal rods. Two types of fins are present, two unpaired median fins - the
dorsal fin and the anal fin, and two pairs of paired fins the pectoral and pelvic fins.
In addition there is a caudal fin (heterocercal caudal fin in the case of the Indian
dog-fish).
As understood, the dorsal fin, more of less triangular in shape, is present me-
dially on the dorsal side of the fish. There are two dorsal fins, the first dorsal fin
anteriorly about in the middle of the body and the second dorsal fin posteriorly
midway between the first dorsal fin and the tip of the tail. The median anal fin is
found ventrally behind the cloacal aperture.
The caudal fin has two unequal halves. The upper (dorsal half) is longer (ver-
tebra extends into it), on the ventral side of this lobe is a notch, which divides this
lobe into a short anterior half and a long posterior part.
The lateral fins the pectoral and pelvic are homologous with the fore- and hind-
limbs of tetrapods. They are also triangular in form, the pectoral fin is ventro-lateral
a little behind the head and the pelvic fins are ventral arising in the junction of the
trunk and the tail and enclose the cloacal aperture between them. In male sharks,
there is a cartilaginous rod at the base of the pelvic fins called as the claspers with
a groove on its inner side.

2.4 Digestive system

Scoliodon sorrokowah’s has a digestive system comprising of the following parts: 1)
buccal cavity; 2) pharynx; 3) oesophagus; 4) stomach; 5) intestine and 6) rectum.
They have teeth similar in structure to the placoid scales, larger and arranged in
several rows at the edges of the jaws. These are modified scales, present in the
mouth, for feeding.
Mouth opens into a buccal cavity that is bordered with teeth. The cavity does
not have any internal nostrils (nostrils do not open internally but are just blind
sacs facilitating olfaction). They have a non-muscular and immobile tongue.
The buccal cavity leads to the pharynx, which is perforated with gill slits (ie, have
branchial apertures on the sides).
The oesophagus is a short region following the pharynx and it leads to a U-
shaped stomach. The stomach consists of two parts: a) the cardiac stomach,
which runs to the hind end of the body cavity; and b) the pyloric stomach that
runs forwards, so lies by the side of the cardiac stomach, thus imparting a U or J
shape to the stomach. At the junction of the pyloric and cardiac stomach, there is
a blind sac that we call as the caecum.
The intestine follows the stomach and is very short (compared to other verte-
brates) and not coiled, it is conspicuous by possessing the spiral valve or the scroll
valve. Spiral valve is a cork-screw shaped structure, running two or three times
around the lower part of the intestine and is involved in increasing the surface area
of the intestine so that the shortness of the intestine is compensated for. At the
junction between the pyloric stomach and the intestine is the pyloric constriction,

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8 2.5. RESPIRATION

followed by a muscular part of the intestine termed as the duodenum or the bursa
entiana. This is followed by the ileum or the posterior region where we find the
spiral valve.
Following the ileum is the rectum which is the final portion of the alimentary
canal which opens to the exterior through the cloaca. The rectum is connected
with teh caecal glands, the urinogenital ducts also open into the cloaca.

2.5 Respiration
Respiration in shark is effected by gills. As said above, the nostrils are blind sacs,
ending blindly supplied by the olfactory nerves, which only possess the sensory
function (thus nose/nostrils have nothing to do with respiration).
The pharynx at its side have five pairs of gill pouches, separated from one
another by interbranchial septa. The pouch opens to the, interior (pharynx)
through the internal branchial aperture and outside (water) through the external
branchial aperture. The walls of the pouches, have mucous membrane lining it which
gives off numerous horizontal folds called gills (or gill filaments to be specific).
Gills are borne on the posterior side of the hyoid arch and on both sides of the
first four branchial arches (there are nine hemibranchs), each of the four branchial
arches possess gill filaments on both sides, that is hemibranchs on both sides, such
an arrangement is called as holobranch or this is a full gill.
Towards the inner side (pharyngeal side) of the branchial arches are comb like
projections called gill rakers, which prevent food from entering the gill slits and
direct the food to the oesophagus.
Water is sucked into the mouth by opening the mouth, next the mouth is
closed and the cavity of pharynx is expanded, water from the buccal cavity passes
through the gill slits outwards, bathing the gills. Water and blood supplied to
the gill filaments exchange gas by counter-current exchange mechanism. Some
elasmobranchs have another mechanism (similar to tunas) to keep their mouth open
while swimming so that water moves into the buccal cavity and pass out through
the gill slits bathing the gills and thus facilitating gas exchange, such a method of
respiration is called as ram ventilation. Most of the sharks have to keep moving
to survive since they do not have an operculum so cannot pump water. Spiracles
present on both sides of the shark just behind the eyes, are openings that lead water
inside to the pharynx, in the process supplying oxygen exchange directly to the eyes
and is an alternate route of respiration.

2.6 Circulatory system

Circulatory system in sharks consists of an ’S’ shaped heart. With two distinct
chambers named as the auricle (or atrium) and the ventricle. The sinus veno-
sus and the conus arteriosus along with the atrium and ventricle makes up the
structures of the heart. Valves are present at the junction between these four struc-
tures. In the conus arteriosus there are several rows of valves. All the valves have
a function to regulate the blood flow in a single direction. These chambers are
contractile and the contraction usually starts with the sinus venosus. The RBC’s in
sharks are nucleated.

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CHAPTER 2. SUPER-CLASS PISCES 9

Paired dorsal aortae Internal

carotid
Celiac artery artery

Efferent branchial
artery V IV III II I FIGURE 12.15 Aortic
VI
Stapedial
of a shark. (a) Embryonic modi
Pretrematic branch artery of aortic arches.New additions (
Posttrematic branch
Efferent and the arches establish the pretrema
Afferent branchial artery
afferent spiracular posttrematic parts of the collecti
Sinus
venosus arteries that receive afferent and supply e
Atrium
h12_451-502.qxd 12/23/10 5:05 PM Page 460 branchial arteries,derivatives of v
and dorsal sections of the aortic
External carotid artery respectively.(b) Adult shark aorti
Ventricle derivatives.Roman numerals indi
Coronary Commissural artery Ventral aorta
artery aortic arches.
Conus arteriosus
(b) After Kent.

verymesenteric
Posterior little mixing of
artery oxygenated and
Anterior deoxygenated
mesenteric artery blood The short section of dorsal aorta between aort
actually occurs, thanks largely, as we see later in this chap- III and IV, termed thecarotidduct, usually closesat m
Internal carotid artery
ter, to the role played by the partially divided heart. phosis. This forces the carotids to fill with blood from
Paired dorsal
Genital artery aortae ativeof theventral aorta. Thesection of ventral aorta
AmphibiansRenal The same mistakenUnpaired
view that the cardiovascu-
Celiac artery arches III and IV becomes thecommon carotid arte
Caudal larartery artery dorsal aorta
system was imperfectly designed was also held about feedstheAortic arches
external carotid (fromtheanterior ventral a
amphibians and for much the same reasons. The anatomical the internal carotid (the anterior section of the do
arrangement of their aortic archessuggeststhat somemixing together with the third aortic arch). The carotid
occurs between oxygenated blood from the gills and deoxy- small cluster of sensory cellsassociated with capillarie
Renal
portal genated blood returning from the body. located near the point at which the common carotid
vein In amphibians, the first two aortic arches (I, II) disap- Its functions are not completely known. Certainly th
pear early in development. The pattern of the remaining body plays a role in sensing the gExternal
as content or pressu
Iliac arches differs between larvae and metamorphosed adults. In blood as well ashavingsome endocrine carotid functions.
Lateral Ventral
arterymost larval salamanders, thenext threeaortic arches(III–V)
Hepatic The next two arches(IV, V)
aorta constitutemajor
artery
abdominal
carry external gills, and the lastportal
vein
aortic arch (VI) sprouts the vessels that join the dorsal aorta. The final aortic a
pulmonary artery to the developing lung. A notable excep- also joins the dorsal aorta, its last short section for
tion is the neotenic salamander Necturus , in which part ofSubclavian
Subclavian ductusarteriosus. Shortly beforejoiningthedorsal a
thesixth arch disappearsand onlyitsdorsal artery
section persists,vein sixth aortic arch givesoff thepulmonaryartery, wh

Figure 2.1: Circulatory system in shark is shown, the figure in the upper half shows
the aortic arches and the heart, the lower panel shows the circulatory system with
dark vessels being the venous system.

From the conus arteriosus continues anteriorly (towards the head region) the
ventral aorta, which gives off four pairs of vessels. The first vessel divides into two
(immediately) so that in total there are five vessels. These five vessels supply blood
to the gill arches or gills and are known as the afferent branchial arteries, they
split into smaller capillaries in the gills.
The blood carried towards the gills are oxygenated and taken back by the ef-
ferent branchial arteries. There are nine efferent arteries and the first eight of
them (one pair each on both sides) join to form four epibranchial arteries on each
side (so four pairs) these four pairs of epibranchial arteries join to form the dorsal
aorta. The dorsal aorta runs backward, giving off branches to various organs. The
ninth efferent artery is connected to the one immediately preceding it. The affer-
ent branchial artery and the corresponding efferent branchial artery constitute an
arterial arch.
The dorsal aorta gives off many branches, the subclavian to the pectoral fins,
the coeliaco-mesentric dividing into two as the coeliac supplying the stomach
and liver, and as the anterior mesentric supplying the intestine and the pancreas.
The lienogastric artery supplies the spleen, intestine and the stomach, the parietal
arteries from which four pairs of renal arteries arise, the posterior mesentric artery
to the rectum and a pair of iliac arteries to the pelvic fins.
The venous system consists of thin (walled) veins; the larger veins are called

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10 2.7. URINOGENITAL SYSTEM

as sinuses and are much distinguished. The veins from different parts of the body
empty into the sinus venosus. Two hepatic portal veins bring back blood from the
liver into the sinus venosus. There are two Sinus Cuvieri or Cuvierian ducts, one on
each side, each Cuvierian duct is formed by the union of the anterior cardinal sinus
and the posterior cardinal sinus; the anterior cardinal sinus collects blood from the
dorsal part of the head, and the posterior cardinal sinus is joined by different sinuses.
From the ventral region of the head the inferior jugular vein drains blood and joins
the sinus venosus close to the joining point of the Cuvierian ducts. The posterior
cardinal sinus of two (lateral) sides join together near the kidneys, the renal veins
drain into this posterior cardinal sinus. Anteriorly each posterior cardinal sinus is
drained by, branchial vein from the pectoral fin, and the iliac vein collects blood
from the pelvic fin and empty into the lateral abdominal vein. From the caudal
region (tail) the veins pass into the kidney as two branches and there in the kidney
split into capillaries and form the renal portal system. The lateral abdominal
vein, on two sides start from behind the kidneys and empty into the posterior
cardinal sinus near the sinus venosus. There is a hepatic portal system that is
drained from the alimentary tract, pancreas, and the spleen, these enters the liver
and break up into capillaries forming the hepatic portal circulation. This empty into
the sinus venosus through the hepatic sinuses.

2.7 Urinogenital system

2.7.1 Excretory organs
A pair of kidneys are present which are mesonephric[1 ] in origin. Which are ribbon
shaped and placed from the base of the gills to the front of the cloaca. Each
kidney has an anterior and posterior part, the anterior part is functionless and
almost atrophied in the females while it acts as the genital canal in the males;
the posterior part has the excretory function, and is the functional kidney in
both males and females. The pronephros is embryonic and disappears and which
paves way for the mesonephros (which is the functional kidney) in adults. The
pronephric duct becomes divided longitudinally into two, one of which is connected
with the mesonephric tubules and known as the mesonephric duct also known
as the wolffian duct, the posterior or second longitudinal tube develops as the
mullerian duct in the females and is atrophied in the males.
The wolffian duct acts as the urinogenital duct in males and just as the urinary
duct in females. In females the mullerian duct function as the genital duct. In
males the urinary ducts of both sides open separately into the cloaca (from both
sides somewhat antero-laterally), the urinary ducts are formed by the fusion of each
urinary tubules from the mesonephros. Some consider this as the ureter (which is
somewhat a tentative consideration).
1 Kidneys develop in the intermediate mesoderm of the dorsal posterior body wall. At the

onset of differentiation first the nephric ridge (an expansion of the mesoderm) is formed which is
slightly protruding from the dorsal body wall. Later in the development develops the nephrotomes
which are segmental in nature and function as primitive kidneys in embryos. It has glomerulus,
bowmans capsule, convoluted tubule and empty into the nephric duct. Developmental and struc-
tural differences in the nephric tubules that arise within the nephric ridge inspired a view of kidney
formation known as the tripartite concept. This concept envisions formation of nephric tubules
in one of three regions in the nephric ridge, the cranial end (anterior) portion being called the
pronephros, the middle part being called as the mesonephros and the posterior part being called
as the metanephros.

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CHAPTER 2. SUPER-CLASS PISCES 11

Urinogenital system of Elasmobranchs

Funnel
Funnel of
rudimentary Ovary
oviduct

Testis Testis Oviduct

Epididymis
Archinephric Kidney
duct (ductus
deferens) Testis
Ovisac
Kidney

Uriniferous Sperm Kidney

kidney duct
Tubular Archineph
Archinephric
duct
Urinary portion of duct
duct testis Urinary
c bladder
Archinephric Urinary
duct duct In
Urogenital
Intestine papilla
Cloaca Cloaca FIGURE 14.23Cloaca Fish ovaries. (a) Hagfish

Male Female
the ovary of the guppy Poecilia reticulata.Ova are fe
development.There can be one to seven oocytes in
(a)
embryos are illustrated.
(a) After Hardisty; (b) after Lambert.

Urinogenital ducts of Elasmobranchs

Chapter Fourteen Funnel

Epididymis
Shell
gland
Ovary

Testis

..
Mullerian
duct
(regressing) Testicular Testicular
Isthmus of
duct duct oviduct
..
(Mullerian
Accessory Archinephric duct)
Archinephric
Archinephric
urinary Archinephric duct
duct duct
duct (vas
duct (opisthonephric duct)
(opisthonephric (opisthonephric
deferens)
duct) duct) Uterus

(a) Shark (b) (a) Shark (b)

(d) Teleost
Male Female
Figure 2.2: Urinogenital system in elasmobranchs: In the above panel we see the
complete urinogenital system of a male and a female torpedo (sting ray), in the lower
panel we see the oviducts and urinogenital ducts in females and males respectively
in a side view showing only one half.

In females, the wolffian duct is continued towards posteriorly (behind/end) as

the urinary ducts, into which each and every urinary tubules open. The ducts unite

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12 2.8. NERVOUS SYSTEM

and open behind by a single aperture into the urinary sinus. So wolffian duct has
genital function in males while act as urinary duct in females, in males a separate
urinary duct (ureter) is formed by the fusion of mesonephric tubules.

2.7.2 Genital organs

In males there are two testes (soft, elongate, lobular bodies) attached to the
dorsal wall of the body by peritoneal folds. In the anterior region of the kidney
(head region) the tubules (urinary tubules otherwise) loose their excretory function
and become connected to the testis. The testis has outgrowth which connect to the
tubules forming the vasa efferentia. The mesonephric duct is well developed and
thrown into coils forming the vas deferens a duct for the passage of the sperms
to the exterior. The vas deferens dilates towards the posterior part forming the
seminal vesicle, each of these from both sides open into the urinogenital sinus
independently which then empties into the cloaca. Two sperm sacs are also present
connected to the urinogenital sinus. Two openings leads from the cloacal opening
to the claspers.
In females, one pair of ovaries are suspended by folds of the peritoneum. The
paired oviducts, also called as the Mullerian duct, converge to the front side and
open into the body cavity by a common opening. They are not connected to the
ovaries in any way. Just above midway there is an enlargement of the mullerian
duct which is called as the shell gland. Behind this shell gland the tube dilates and
is called as the uteri. The uteri join together and form a vagina which opens into
the cloaca.
Sperms pass from testes to vas efferentia to the vas deferens, and finally to the
seminal vesicles here the sperm is stored. During copulation claspers are inserted into
the cloaca of the female and sperms pass into the female. The sperm sacs function
as organs that force the movement of sperm from the clasper to the cloaca. Ova
when ripe fall into the body cavity and pass into the oviducts. Fertilization occurs
somewhere in the anterior part of the oviduct, above the shell gland.
Epigonal organs are closely associated with the caudal part of the gonads of
sharks. These are lymphoid bodies which are centres of formation of erythrocytes
and leucocytes.

2.8 Nervous System

The Nervous system can be divided into two, the Central Nervous System (CNS)
and the Peripheral Nervous System (PNS). The PNS comprises all the nervous
tissue outside the CNS, while the CNS comprises the brain and the spinal cord. The
nervous system receives impulses from various receptors and transmits signals to
various effectors. Effectors can be mechanical in nature - muscles, or chemical in
nature such as glands, which result in muscle contraction and glandular secretions.
Two types of cells make up the nervous system, the neurons and the neuroglial
cells or the glia. The neuroglial cells do not transmit impulses, but they support,
nourish and insulate the neurons. Microglia, engulf foreign material and bacteria
(protection function), and oligodendroglia and schwann cells insulate the axons.
The neuroglial cell that forms the mylein sheath in the CNS is the oligodendroglial
cell and those that form the mylein sheath in the PNS are the schwann cells.

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CHAPTER 2. SUPER-CLASS PISCES 13

Neurons are specialised for transmitting the electrical signals through-out the
body, upto long distances. They are the structural and functional units of the
nervous system. As we already know they have a cell body, one or more dendrites
and one axon in most cases. There are unipolar, multipolar and bipolar neurons.
There are certain neurosecretory cells that have endocrine function. Normal
neurons produce neuro-transmitters at the synapses, and they help in transmis-
sion of the impulses, the neurosecretory cells also produce neurotransmitters at the
synapses, but they are secreted into the capillaries and are transmitted to the target
tissues.

2.8.1 Peripheral nervous system

Peripheral nerves serve either somatic or visceral tissues and carry sensory or motor
information. Somatic nerves pass to or from somatic tissues: skeletal muscle, skin,
and their derivatives. Visceral nerves pass to or from viscera: involuntary muscles
and glands. Nerves carrying information from tissues to the CNS are afferent, or
sensory, neurons. Nerves carrying information away from the CNS to effectors are
efferent, or motor, neurons. Thus, a somatic sensory nerve might carry information
about touch, pain, or temperature from the skin to the central nervous system. A
somatic motor nerve carries impulses from the CNS to a striated muscle to stimulate
its contraction. A visceral sensory nerve delivers information about the condition
of internal viscera to the CNS. A visceral motor nerve innervates visceral effectors
(cardiac muscle, smooth muscle, or glands). The components of the PNS that
control visceral activity constitute the autonomic nervous system (ANS).
There are two functional divisions for the autonomic nervous system (ANS), the
sympathetic system and the parasympathetic system. The ANS controls the
visceral activity by these two antagonistic nervous systems.
The sympathetic nervous system prepares the body for strenuous action by
increasing activity of the viscera, although it slows digestive processes. Stimula-
tion of the sympathetic system inhibits activity of the alimentary canal but promotes
contraction of the spleen (causing it to release extra red blood cells into the general
circulation), increases heart rate and blood pressure, dilates coronary blood vessels,
and mobilizes glucose from glycogen storage in the liver. It is often said that the
sympathetic nervous system prepares the individual to fight or flee.
The parasympathetic nervous system restores the body to a restful or vege-
tative state by lowering its activity level, although digestion is stimulated. The
effects of the parasympathetic system are antagonistic to those of the sympathetic
system. It enhances digestion, slows heart rate, drops blood pressure, constricts
coronary vessels, and promotes glycogen formation.
Adrenergic and Cholinergic Control: The sympathetic system is said to be
adrenergic because the neurotransmitters released during stimulation are adrenaline
or noradrenaline (also termed epinephrine and norepinephrine). The parasympa-
thetic system is said to be cholinergic because the neurotransmitter released is
acetylcholine.

2.8.2 Central Nervous System

The central nervous system primarily coordinates activities that enable an organism
to survive and reproduce in its environment.

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14 2.8. NERVOUS SYSTEM

Spinal cord
Spinal cord is the part of the CNS, and has two parts named so due to their
appearance in fresh preparations, the grey matter and the white matter which is
external. The grey matter is crescent shaped in each half and is found in the core
region, thus there are two horns for the grey matter of one half. From the spinal
cord arises two nerves one in the dorsal side and another in the ventral side, of each
half. The dorsal root gives rise to a ganglion shortly after its emergence and then
continues forward, it meets with the ventral root subsequently, and forms the spinal
nerve. The vertebral column has a segmental appearance and each segment has
one pair of roots originating (in both sides). Thus each spinal nerve provides to its
adjacent dermatome maintaining the segmental nature.

Figure 2.3: Figure of the spinal cord in a cross section, showing the spinal nerves.

The spinal nerve immediately after its origin (at the joining of the two roots)
gives rise to a ramus communicans which inturn runs towards the ventral side and
joins with the sympathetic chain which is parallel to the spinal cord. The ventral
root has the motor neurons and the dorsal root has the motor neurons. These two
roots are connected internally in the grey matter by neurons. The peripheral spinal
nerve is of mixed nature. One sensory neuron originates from the dorsal ganglion
and has two arms, one arm towards the exterior and one arm towards the spinal
cord. The branch into the spinal cord takes the impulses towards the CNS (brain).

Brain
Embryologically the brain forms as a wide region in the anterior of the (embryonic)
neural tube. The posterior part is the hind brain or rhombencephalon (divided
into metencephalon and the mylencephalon) which forms the medulla oblongata,
cerebellum and the pons, then immediately in front of the hind brain is the mid-
brain or mesencephalon which has a sensory tectum and a motor tegumentum,
then the anterior part of the brain is the fore brain or prosencephalon which has
cerebrum or telencephalon, and the thalamus or the diencephalon.
The Prosencephalon (or cerebrum) has two lateral ventricles (spaces) internally,
and anterior to it is the olfactory lobes, which should be most probably associated

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CHAPTER 2. SUPER-CLASS PISCES 15

Telencephalon

Diencephalon

Mesencephalon

Metencephalon

Myelencephalon

Figure 2.4: Brain of the shark in (left) dorsal and (right) ventral views, with regions
of the brain color-coded.

with the well developed sense of smell. The olfactory lobes are connected to the
cerebrum by long olfactory peduncles. The lateral ventricles in the cerebrum contin-
ues into the olfactory lobes as olfactory ventricles. This makes up the telencephalon,
the diencephalon is short and hidden from view by the forward prolongation of the
cerebellum (hind brain). The diencephalon has four distinct regions, the dorsal tha-
lamus, the ventral thalamus, the epithalamus and the hypothalamus. The floor of
diencephalon bears the infundibulum and the pituitary body. At the sides of the
infundibulum are two thin-walled oval sacs, the lobi inferiores which are produced
behind into two vascular expansions called the sacci vasculosi. The non-vascular
roof bears the pineal body on a long stalk extending forwards (epithalamus). The
infundibulum is an outgrowth of the hypothalamus and it fuses with an outgrowth of
the stomodaeum and the two together forms the pituitary body (or hypophysis). It
consists of an anterior (adenohypophysis) and a posterior (neurohypophysis) parts.
The mesencephalon consists of two optic lobes hidden dorsally by the cerebrum
and ventrally by the infundibulum. On the ventral side in the region of the optic
lobes are the crura cerebri.
The metencephalon (anterior half of the rhombencephalon), or the cerebellum
is very large and extends in front and behind overlapping optic lobes and medulla
oblongata. The medulla oblongata is produced anteriorly on the dorso-lateral sides
into two ear lappets, the corpora restiformia.
There are ten pairs of cranial nerves, the origin and the positions shown in the
figure below. The cranial nerves from one to ten are: Olfactory (S), Optic (S),
Occulomotor (Mo), Trochlear (Mo), Trigeminal (S/M), Abducens (M), Facial (M),

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16 2.9. ACCESSORY RESPIRATION IN FISHES

Auditory (S), Glossopharyngeal (S) and Vagus (M). [M = mixed; Mo = motor; S

= sensory].

Trochlear n. (IV) Optic lobe

Cerebellum
Auricle of
Epiphysis cerebellum Trigeminal (V) & preotic
lateral line nn.
Cerebral hemisphere Vagus (X) & postotic
lateral line nn.
Terminal n. (0)
Occipital nn.
Olfactory sac Glossopharyngeal n. (IX)
Statoacoustic n. (VIII)
Olfactory bulb Optic n. (II) Facial n. (VII)
Olfactory tract Optic chiasm Abducens n. (VI)
Inferior lobe Oculomotor n. (III)
Vascular sac

Figure 2.5: Brain and cranial nerves of the shark in left lateral view.

2.9 Accessory respiration in fishes

In all tropical regions of the world, deoxygenation is frequent in shallow and stagnant
waters.Whenever water is slow-moving, covered with vegetation, and heavily shaded
so that photosynthesis is slight, and where cooling at night is not sufficient to
produce overturn, deoxygenation is bound to occur. The only oxygen in these
waters gets there by diffusion from the air.
Usually freshwater habitats are the only ones effected, but also estuarine habitats
are susceptible, especially mangrove swamps. Many bony fishes inhabiting such
waters have evolved special air-breathing structures to cope with these unusual
conditions. Others have evolved ways to live out of the water for extended periods
of time.
Air-breathing capability has evolved independently in many relatively unrelated
groups of fishes, in preteleosts as well as teleosts.
The accessory air-breathing structures of bony fishes are all modifications of the
epithelium of the alimentary canal, or gill chamber, or diverticula emerging from
these parts. This includes a host of structures: the mouth, pharynx, gill chamber,
esophagus, airbladder, stomach, and intestine.
All have in common a specialized epithelium that is often equipped with branched,
tree-like outgrowths to increase the surface area. Lying underneath this are a rich
network of blood capillaries. Gills are never used for air breathing because the gill
lamellae are not stiff enough to support themselves and collapse when out of water.
The surface area is lost and the fish suffocates.
All air-breathing organs function just as absorbers of oxygen, they never have
anything to do with carbon dioxide. The gills and/or the skin take care of the excre-
tion of carbon dioxide. Most air-breathers have evolved a new organ by modifying
some existing structure. This new organ deals with oxygen uptake from the air.
Elimination of carbon dioxide is done by way of the gills and in some cases scales
are reduced as well and the skin is used for the discharge of carbon dioxide. Swamp
eels (Synbranchiformes) and the famous mudskipper (genus Periophthalmus) fall
into this latter category (of less scales).
The air from outside is drawn into these (additional/new) chambers by the
mouth and retained there for the aeration of the blood. Anabas, Amphipnous,

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CHAPTER 2. SUPER-CLASS PISCES 17

Channa, Clarias, Saccobranchus etc. are some Actinopterygian fishes with such
organs.
Anabas has the habit of migrating from pond to pond and during this period
of progression on land, the fish breathes air from the atmosphere. There are two
air chambers, one on each side of the head. These are extensions of the branchial
cavities. They contain concentrically arranged wavy plates (labrynthine organs)
which are outgrowths of the upper part of the first branchial arch. The plates are
covered with vascular membrane. Each chamber communicates with the pharynx by
the first gill-slit, and with the gill-cavity by an opening situated between the hyoid
and the first branchial arch. Air is drawn in by the mouth into the air chamber by
the first gill-slit, and it passes out by the branchial aperture. The fish, it is said,
can live for six or seven hours out of water.
In Amphipnous, the air chambers are saccular outgrowths of the dorsal walls
of the pharynx, extending as far as the third branchial arch. The walls of the sacs
are folded and vascular. The sacs communicate with the pharynx by an opening
through which air is drawn in, and the air is forced out through the gill slits and
thence by the opercular opening. Here the gills are reduced to a few filaments in
the first two arches.
In Channa, a pair of air chambers are present on each side of the head, arising
as outgrowths of the pharynx, above the first gill arch and extending as far as the
last gill cleft. Processes of the hyomandibular arch, and the epibranchials of the
first two branchial arches project into the chamber which are vascular. Air passes
to these chambers through the mouth and out by the opercular aperture.
The catfishes of claridae, have air chambers which are outgrowths of the gill
cavities and they enclose branched vascular processes (arborescent or dendriform
organs). There are two such organs which are outgrowths of the upper parts of the
second and fourth branchial arches. In Saccobranchus there are a pair of thubular
sacs as outgrowths of the gill chambers extending as far as the middle of the tail
region. At the hind end of the tube, there are folds forming a sort of air chamber.
THe air chamber communicates with the buccal cavity by a slit through which air
passes in and out.
Periophthalmus (mudskipper) is an esturine fish, common in brackish waters,
skipping about on the mudflats of the mangroove swamps. In this fish, the large
opercular cavities are filled with air drawn through the mouth. It is said that the
mudskipper has become so accustomed to life out of water that it dies of suffocation
if prevented from coming out for prolonged periods. In Andamia (Blennidae) a
tropical marine fish, popularly known as the rock skipper, the opercular chambers
serve aerial respiration.

2.10 Classification of Fishes

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18 2.10. CLASSIFICATION OF FISHES

Table 2.1: Classification of Chondrichthyes (In bold are the Orders, and in italicz
are the families each of them are followed by the common names)
Class Chondrichthyes Total car-
tilaginous fishes 188 (genera)
1,168 (species)
Order Family

(Subclass Holocephali)

Chimaeriformes Modern chi-

maeras (Total chimaeras 6 gen-
era 43 species)
Callorhinchidae Elephant fishes
Rhinochimaeridae Longnose chi-
maeras
Chimaeridae Shortnose chi-
maeras

(Subclass Elasmobranchii) (To-

tal elasmobranchs 182 genera
1,125 species)

Superorder Squalomorphii
Hexanchiformes Cow and frilled
sharks
Chlamydoselachidae Frilled
sharks
Hexanchidae Sixgill and Sev-
engill sharks
Squaliformes Dogfish sharks
Echinorhinidae Bramble sharks
Squalidae Dogfish sharks
Centrophoridae Gulper sharks
Etmopteridae Lantern sharks
Somniosidae Sleeper sharks
Oxynotidae Roughsharks
Dalatiidae Kitefin sharks
Squatiniformes Squatinidae Angel sharks
Pristiophoriformes Sawsharks Pristiophoridae

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CHAPTER 2. SUPER-CLASS PISCES 19

Table 2.2: Classification of Chondrichthyes (continued..)

Class Chondrichthyes
Order Family

Rajiformes Batoids (rays)

Suborder Pristoidei:
Sawfishes
Suborder Rhinoidei:
Sharkrays
Suborder Rhynchoba-
toidei: Wedge fishes
Suborder Rhinobatoidei:
Guitarfishes
Suborder Platyrhinoidei:
Thornbacks
Suborder Zanobatoidei:
Panrays
Suborder Torpedinoidei:
Electric rays
Narcinidae Numbfishes
Narkidae Sleeper rays
Hypnidae Coffin rays
Torpedinidae Torpedo rays
Suborder Rajoidei:
Skates
Arhynchobatidae Softnose
skates
Rajidae Hardnose skates
Anacanthobatidae Legskates
Suborder Myliobatoidei:
Stingrays
Plesiobatididae Giant stingarees
Urolophidae Stingarees
Urotrygonidae Round stingrays
Hexatrygonidae Sixgill stingrays
Potamotrygonidae River
stingrays
Dasyatidae Whiptail stingrays
Gymnuridae Butterfly rays
Myliobatidae Eagle rays
Rhinopteridae Cownose rays
Mobulidae Devil rays

Superorder Galeomorphii

Heterodontiformes Bullhead Heterodontidae

sharks

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20 2.10. CLASSIFICATION OF FISHES

Table 2.3: Classification of Chondrichthyes (continued...

Class Chondrichthyes
Order Family

Orectolobiformes Carpet sharks

Parascylliidae Collared carpet-
sharks
Brachaeluridae Blind sharks
Orectolobidae Wobbegongs
Hemiscylliidae Longtailed car-
petsharks
Ginglymostomatidae Nurse
sharks
Stegostomatidae Zebra sharks
Rhincodontidae Whale sharks
Lamniformes Mackerel sharks
Odontaspididae Sand tiger
sharks
Pseudocarchariidae Crocodile
sharks
Mitsukurinidae Goblin sharks
Megachasmidae Megamouth
sharks
Alopiidae Thresher sharks
Cetorhinidae Basking sharks
Lamnidae Mackerel sharks
Carcharhiniformes Ground
sharks
Scyliorhinidae Catsharks
Proscylliidae Finback catsharks
Pseudotriakidae False catsharks
Leptochariidae Barbeled hound-
sharks
Triakidae Houndsharks
Hemigaleidae Weasel sharks
Carcharhinidae Requiem sharks
Sphyrnidae Hammerhead sharks

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CHAPTER 2. SUPER-CLASS PISCES 21

Figure 2.6: The classification of sharks as a figure.

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22 2.10. CLASSIFICATION OF FISHES

Table 2.4: Classification of Actinopterygii the ray finned fishes

Class (Osteichthyes) Actinopterygii

Subclass Chondrostei
Order Polypteriformes, bichirs and reedfishes
Order Acipenseriformes, sturgeons and paddle-
fishes
Subclass Neopterygii
Infraclass Holostei
Order Lepisosteiformes, gars
Order Amiiformes, bowfins
Infraclass Teleostei
Superorder Osteoglossomorpha
Order Osteoglossiformes, bony-tongued fishes
Order Hiodontiformes, mooneye and goldeye
Superorder Elopomorpha
Order Elopiformes, ladyfishes and tarpon
Order Albuliformes, bonefishes
Order Notacanthiformes, halosaurs and spiny
eels
Order Anguilliformes, true eels
Order Saccopharyngiformes, gulper eel
Superorder Clupeomorpha
Order Clupeiformes, herrings and anchovies
Superorder Ostariophysi
Order Gonorynchiformes, milkfishes
Order Cypriniformes, barbs, carp, danios, gold-
fishes, loaches, minnows, rasboras
Order Characiformes, characins, pencilfishes,
hatchetfishes, piranhas, tetras, dourado / golden
(genus Salminus) and pacu
Order Gymnotiformes, electric eels and knife-
fishes
Order Siluriformes, catfishes
Superorder Protacanthopterygii
Order Argentiniformes, barreleyes and slickheads
(formerly in Osmeriformes)
Order Salmoniformes, salmon and trout
Order Esociformes pike
Order Osmeriformes, smelts and galaxiids

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CHAPTER 2. SUPER-CLASS PISCES 23

Table 2.5: Classification of Actinopterygii the ray finned fishes (Continued...)

Class (Osteichthyes) Actinopterygii

Superorder Stenopterygii (may belong in Pro-

tacanthopterygii)
Order Ateleopodiformes, jellynose fish
Order Stomiiformes, bristlemouths and marine
hatchetfishes
Superorder Cyclosquamata (may belong in Pro-
tacanthopterygii)
Order Aulopiformes, Bombay duck and
lancetfishes
Superorder Scopelomorpha
Order Myctophiformes, lanternfishes
Superorder Lampridiomorpha
Order Lampriformes, oarfish, opah and ribbon-
fishes
Superorder Polymyxiomorpha
Order Polymixiiformes, beardfishes
Superorder Paracanthopterygii
Order Percopsiformes, cavefishes and trout-
perches
Order Batrachoidiformes, toadfishes
Order Lophiiformes, anglerfishes
Order Gadiformes, cods
Order Ophidiiformes, pearlfishes
Superorder Acanthopterygii
Order Mugiliformes, mullets
Order Atheriniformes, silversides and rainbow-
fishes
Order Beloniformes, flyingfishes
Order Cetomimiformes, whalefishes
Order Cyprinodontiformes, livebearers, killifishes
Order Stephanoberyciformes, ridgeheads
Order Beryciformes, fangtooths and pinecone-
fishes
Order Zeiformes, dories
Order Gobiesociformes, clingfishes
Order Gasterosteiformes, sticklebacks
Order Syngnathiformes, seahorses and pipefishes
Order Synbranchiformes, swamp eels
Order Tetraodontiformes, filefishes and pufferfish
Order Pleuronectiformes, flatfishes
Order Scorpaeniformes, scorpionfishes and the
sculpins
Order Perciformes, 40% of all fish, including an-
abantids, centrarchids (including bass and sun-
fish), cichlids, gobies, gouramis, mackerel, tuna,
perches, scats, whiting, wrasses, bettas

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24 2.10. CLASSIFICATION OF FISHES

Table 2.6: Classification of Sarcopterygii the lobe finned fishes; the Coelacanth and
the lung fishes.
Class Sarcopterygii

Class Sarcopterygii
Infraclass Dipnomorpha
Subclass Dipnoi
Order Ceratodontiformes, Neocerato-
dus forsteri (also known as the Aus-
tralian lungfish, Burnett salmon, and bar-
ramunda)
Order Lepidosireniformes, Lep-
idosirenidae (the South American
lungfish) and Protopteridae (the African
lungfish)
Infraclass Crossopterygii
Order Actinistia, contains the coela-
canths, including the two living coela-
canths, the West Indian Ocean coela-
canth Latimeria chalumnae, and the king
of the sea Latimeria menadoensis.
Infraclass Tetrapodomorpha
Superclass Tetrapoda

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CHAPTER 2. SUPER-CLASS PISCES 25

2.11 Examples
2.11.1 Selachii
2
Examples of two cartilaginous fishes

Pristis (Order Rajiformes; sub-order Myliobatoidei) Pristis is otherwise called

as the sawfish. Sawfishes, also known as carpenter sharks, are a family (Pristidae)
of rays characterized by a long, narrow, flattened rostrum, or nose extension, lined
with sharp transverse teeth, arranged so as to resemble a saw. Several species of
sawfishes can grow to about 7 m (23 ft). The sawfish’s most distinctive feature is
the saw-like rostrum, covered with electrosensitive pores that allow the sawfish to
detect slight movements of prey hiding in the muddy sea floor. The rostrum also
serves as a digging tool to unearth buried crustaceans. The body and head of a
sawfish are flat, and they spend most of their time lying on the sea floor. Like rays,
a sawfish’s mouth and nostrils are on its flat underside. The mouth is lined with
small, dome-shaped teeth for eating small fish and crustaceans; sometimes the fish
swallows them whole. Sawfish breathe with two spiracles just behind the eyes that
draw water to the gills. The skin is covered with tiny dermal denticles that gives
the fish a rough texture. Like other elasmobranchs, sawfish lack a swim bladder and
use a large, oil-filled liver to control buoyancy.
All species of sawfishes are listed as Endangered or Critically Endangered by the
IUCN, and face the threat of extinction as a result of habitat loss and overfish-
ing. Sawfishes are marine, euryhaline (moving between freshwater and saltwater),
or marginal (brackish water) species, and are widely distributed across tropical and
warm temperate nearshore ocean waters in the Atlantic and Indo-Pacific. Sawfishes
are nocturnal, usually sleeping during the day and hunting at night. Despite fear-
some appearances, they do not attack people unless provoked or surprised. The
small-tooth sawfish is well known by fishermen as a prize game fish because of the
fight it puts up once hooked.

Trygon (Order Rajiformes; sub-order Pristoidei) The fishes of this genera are
generally called as sting rays. Body flattened with pectoral fins greatly expanded
and fused with head and trunk; tail slender or whip-like, usually with one or several
spines; usually with a single dorsal fin, but no caudal fin. No electric organ. Though
not aggressive, the common stingray can inflict an excruciating wound with its ser-
rated, venomous tail spine. This species is not sought after by commercial fisheries,
but is taken in large numbers as ’bycatch’ and utilized for food, fishmeal, and liver
oil. The pectoral fins or "wings" of this species are sold smoked or dried and salted,
and it is also utilized as a source of fishmeal and liver oil. The populations of the
sting-rays are dwindling and are considered as ’threatened’ by the IUCN.
The fish belongs to the family Dasyatidae and are called as teh whiptail stingrays
belonging to the order Myliobatiformes. They are found worldwide in tropical to
temperate marine waters, and a number of species have also penetrated into fresh
water in North America, Africa, Asia, and Australia. Members of this family have
2 The fishes listed under this subsection are actually the fishes under sub-class Elasmobranchii
according to the most recent classification. Class: Chondrichthyes is divided into Elasmobranchi
and Holocephali (two sub-classes) in the most recent classification. The rays are under the Order
Rajiformes according to the present classification - the Stingray in the sub-order Myliobatoidei
and the sawfish in the sub-order Pristoidei

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26 2.11. EXAMPLES

flattened pectoral fin discs that range from oval to diamond-like in shape. Their
common name comes from their whip-like tails, which are much longer than the
disc and lack dorsal and caudal fins. All whiptail stingrays, except the porcupine
ray (Urogymnus asperrimus), have one or more venomous stings near the base of
the tail, which is used in defense. They range in size from 18 cm (7.1 in) to almost
2 m (6.6 ft) across.

2.11.2 Holocephali
The fishes of this class are generally known as chimeras. They are also known
informally as ghost sharks and ratfish. They are the closest relatives to the sharks
(elasmobranchs) though in evolutionary terms, they branched off from sharks nearly
400 million years ago and have remained isolated ever since. Chimaeras live in
temperate ocean floors down to 2,600 m (8,500 ft) deep, with few occurring at
depths shallower than 200 m (660 ft). Exceptions include the members of the
genus Callorhinchus. Chimaera skeletons are constructed of cartilage. Their skin is
smooth and largely covered by placoid scales, and their color can range from black
to brownish gray. For defense, most chimaeras have a venomous spine located in
front of the dorsal fin.
Chimaeras resemble sharks in some ways: they employ claspers for internal
fertilization of females and they lay eggs with leathery cases. However, unlike
sharks, male chimaeras also have retractable sexual appendages on the forehead (a
type of tentaculum) and in front of the pelvic fins. The females lay eggs in spindle-
shaped, leathery egg cases. They also differ from sharks in that their upper jaws are
fused with their skulls and they have separate anal and urogenital openings. They
lack sharks’ many sharp and replaceable teeth, having instead just three pairs of
large permanent grinding tooth plates. They have gill covers or opercula like bony
fishes.
These fishes are thought to have evolved during the Silurian period, although
they are not well characterised (evolutionarily) for the lack of good fossils. The
group is target for phylogenetic studies to understand more about evolution and one
species Callorhinchus milii have its whole genome sequence in draft stages meaning
that in the coming years much more will be understood about the chimeras.

2.11.3 Actinopterygii
Two examples of the ray finned fishes 3 .

Mugil The flathead mullet, Mugil cephalus, is a mullet of the genus Mugil in the
family Mugilidae, found in coastal tropical and subtropical waters worldwide. Its
length is typically 30 to 75 centimetres (12 to 30 in). They belong to the family
Mugilidae and order of ray-finned fish found worldwide in coastal temperate and
tropical waters, and in some species in fresh water. The family includes about 80
species in 17 genera, although half of the species are in just two genera (Liza and
Mugil). The roe 4 of this mullet is salted, dried, and compressed to make a specialty
3 Given
in the syllabus as ’order Acanthopterygii’ which is under sub-class Actinopterygii and
superorder Acanthopterygii; the Mugil being in the Order Mugiliformes and the Mackerel being
under the Order Perciformes
4 Roe or hard roe is the fully ripe internal egg masses in the ovaries, or the released external

egg masses of fish and certain marine animals, such as shrimp, scallop and sea urchins.

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CHAPTER 2. SUPER-CLASS PISCES 27

food across the world.

Rastrelliger kanagurta The Indian mackerel (Rastrelliger kanagurta) is a species

of mackerel in the scombrid family (family Scombridae) of order Perciformes. It is
commonly found in the Indian and West Pacific oceans, and their surrounding seas.
It is an important food fish and is commonly used in South and South-East Asian
cuisine.
The body of the Indian mackerel is moderately deep, and the head is longer
than the body depth. The maxilla are partly concealed, covered by the lacrimal
bone, but extend till around the hind margin of the eye. hese fish have thin dark
longtitudinal bands on the upper part of the body, which may be golden on fresh
specimens. There is also a black spot on the body near the lower margin of the
pectoral fin. Dorsal fins are yellowish with black tips, while the caudal and pectoral
fins are yellowish. The remaining fins are dusky.
The Indian mackerel is generally found in shallow, coastal waters, where the
surface water temperature is at least 17 ◦ C (63 ◦ F). Adults of this species are
found in coastal bays, harbours and deep lagoons. They are commonly found in
turbid waters rich in plankton.

2.11.4 Sarcopterygii
5
Examples of lobe-finned fishes.

Crossopterygii The coelacanths arose in the Devonian period, radiated some-

what, and reached their evolutionary peak in the Mesozoic era. At the end of the
Mesozoic era they nearly disappeared but left one remarkable surviving species, La-
timeria chalumnae. Since the last coelacanths were believed to have become extinct
70 million years ago, the astonishment of the scientific world may be imagined when
the remains of a coelacanth were found on a dredge off the coast of South Africa
in 1938. An intensive search to locate more specimens was successful off the coast
of the Comoro Islands. There fishermen occasionally catch them at great depths
with hand lines, providing specimens for research. This was believed to be the only
population of Latimeria until 1998, when the scientific world was again surprised
by the capture of a coelacanth, possibly representing a new species, in Indonesia,
10,000 km from the Comoros!
The “modern” marine coelacanth is a descendant of the Devonian freshwater
stock. The tail is of the diphycercal type but possesses a small lobe between the
upper and lower caudal lobes, producing a three-pronged structure. Coelacanths
are a deep metallic blue with irregular white or brassy flecks, providing camouflage
against the dark lava-cave reefs they inhabit. Young are born fully formed after
hatching internally from eggs 9 cm in diameter—the largest among bony fishes.
Living examples of coelacanths Latimeria chalumnae have a deep blue color
which probably camouflages them from prey species; however, the Indonesian species
(L. menadoensis) is brown. They are more closely related to lungfish (see below),
amphibians, reptiles and mammals than to the common ray-finned fishes. They are
found along the coastlines of the Indian Ocean and Indonesia
5 The Class Sarcopterygii is divided into two groups of lobe-finned fishes in infraclass Dipnomor-

pha and infraclass Crossopterygii. The fishes listed below belong to Infraclass Crossopterygii, Order
Actinistia (Latimeria) and Infraclass Dipnomorpha/Subclass Dipnoi (African, South American and
Australian lung fishes)

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28 2.11. EXAMPLES

Dipnoi Lobe-finned fish possess a pair of lungs and a fin structure with obvious
homologies with the tetrapod limb. The seven relict species are the freshwater
lungfish (infraclass Dipnoi) and the marine coelacanth (infraclass Coelocanthimor-
pha; Latimeria). The lobe-finned fishes (Class: Sarcopterygii) are the survivors of
a group once abundant during the Devonian period (420-360 million years ago) of
the Paleozoic (541 to 252.2 million years ago).
These lungfishes are a classic example of disjunct distribution, since three
genera of fishes are found in three distant continents (locations; and obviously they
are fresh water fishes) well separated by vast stretches of oceans, in the South
America, Africa and Australia. This, presence in three continents, means that they
should have been present on the earth (in the Pangea which is around 300 - 200
million years old) well before the ’plate tectonic’ movements that separated the
continents to its current positions (fresh water fishes are not supposed to migrate
through the ocean). Their current distribution has thus been influenced by the
continental drift.
All early sarcopterygians had lungs as well as gills, and a tail of the heterocer-
cal type. However, during the Paleozoic the orientation of the vertebral column
changed so that the tail became symmetrical, with the median dorsal and ventral
fins displaced posteriorly to form one continuous, flexible fin around the tail. This
type of tail is called diphycercal.
The strong, fleshy, paired lobed fins of the sarcopterygians (pectoral and pelvic)
may have been used much like four legs to scuttle along the bottom. They had
powerful jaws and their skin was covered with heavy scales that consisted of a
dentine-like material called cosmine overlaid by a thin enamel.
Circulation consist of heart with a sinus venosus, two atria, a partly divided ven-
tricle, and a conus arteriosus. These fishes have a double circulation with pulmonary
and systemic circuits, they characteristically possess five aortic arches.
Of the three surviving genera of lungfishes, most similar to early forms is Neo-
ceratodus (Gr. neos, new, keratos, horn, odes, form), the living Australian lungfish,
which may attain a length of 1.5 m. This lungfish, unlike its relatives, normally
relies on gill respiration, and cannot survive long out of water. The South American
lungfish Lepidosiren (L. lepidus, pretty, siren, mythical mermaid) and the African
lungfish Protopterus (Gr. protos, first, pteron, wing) can live out of water for long
periods of time.
Protopterus lives in African streams and ponds that may dry during the dry sea-
son, with their mud beds baked hard by the hot tropical sun. The fish burrows down
at the approach of the dry season and secretes a copious slime that is mixed with
mud to form a hard cocoon in which it estivates until the rains return. Surprisingly
little is known about the ecology of the South American lung-fish Lepidosiren.
Protopterus: This genera comprises four air-breathing species in Africa. African
lungfish are more carnivorous than the other types of lungfish, they eat molluscans,
crustaceans and some fish. If these lungfish are prevented from reaching the water’s
surface to breathe they will drown. Like Lepidosiren paradoxa, they can survive for
long periods of drought by aestivation.
Neoceratodus: This species is confined to four rivers in Australia. Unlike other
lungfish, it respires almost exclusively through its gills and only uses its single lung
when stressed. This lungfish is also omnivorous and eats plant material.
Lepidosiren: The South American lungfish must breathe air to survive. It can
aestivate for up to four years during the dry season in a nest tube which it digs in
its swampy habitat. This lungfish is omnivorous and eats plant material.

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CHAPTER 2. SUPER-CLASS PISCES 29

The Australian lungfish Neoceratodus forsteri is the least specialized of three

lungfish genera. The African lungfish Protopterus sp. is best adapted of the three
for remaining dormant in mucous-lined cocoons breathing air during prolonged pe-
riods of drought.

Siby Philip - Zoology